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Zootaxa 3750 (2): 101129 ISSN 1175-5326 (print edition)

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Copyright 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3750.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:F01C254C-7CCA-4669-8C30-BF61180BD675

Adults and larvae of two Leucochrysa (Leucochrysa) species


(Neuroptera: Chrysopidae): descriptions, biological notes, and relationships

CATHERINE A. TAUBER1, FRANCISCO SOSA2, GILBERTO S. ALBUQUERQUE3 & MAURICE J. TAUBER1


1
Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853-2601 and Department of Entomology, University
of California, Davis, CA, USA 95616. E-mail: cat6@cornell.edu
2
Museo Entomolgico "Dr. Jos Manuel Osorio" (UCOB), Universidad Centroccidental "Lisandro Alvarado", Venezuela.
E-mail: fransodu73@hotmail.com
3
Laboratrio de Entomologia e Fitopatologia, CCTA, Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, RJ, Bra-
zil 28013-602. E-mail: gsa@uenf.br

Abstract

This taxonomic study includes: (i) a redescription of Leucochrysa (Leucochrysa) nigrilabris (Banks) ( and ), based on
the type specimen and new material and (ii) images of the Leucochrysa (L.) insularis (Walker) type, adult color polymor-
phism, and genital characters ( and ). For both species, it provides: (iii) descriptions of the larvae, (iv) biological notes,
and (v) geographic records. Using the above information, we compare the two species with each other and with other Leu-
cochrysa (Leucochrysa) species that purportedly are closely related. We conclude: First, the larval features of L. (L.) nig-
rilabris and L. (L.) insularis coincide with those previously described as characteristic of the genus Leucochrysa and its
subgenus Leucochrysa. Second, based on their genitalia ( and ), larval morphology, and biology (e.g., deposition of
eggs in clusters), the two species are distinct, but very closely related. And, third, L. (L.) nigrilabris and L. (L.) insularis
share several characteristics with the Leucochrysa (L.) varia-like species; these include similar adult color polymor-
phisms and aspects of their larval morphology. However, their genitalia ( and ) differ significantly from those of the
described L. (L.) varia-like species, and thus we consider the two sets of species to be distinct.

Key words: Chrysopinae, Leucochrysa (L.) nigrilabris, Leucochrysa (L.) insularis, Leucochrysa (L.) varia, oviposition,
color morphs

Resumo

O presente estudo taxonmico inclui: (i) redescrio de Leucochrysa (Leucochrysa) nigrilabris (Banks) ( e ), com base
no espcime-tipo e novo material, e (ii) imagens do tipo, polimorfismo de colorao dos adultos e genitlia ( e ) de
Leucochrysa (L.) insularis (Walker). Para ambas as espcies, apresentam-se ainda: (iii) descries das larvas, (iv) infor-
maes biolgicas e (v) registros geogrficos. Tais informaes so usadas para comparar as duas espcies entre si e com
outras espcies de Leucochrysa (Leucochrysa) que supostamente so proximamente relacionadas. A partir dessas compa-
raes, conclui-se que (1) as caractersticas das larvas de L. (L.) nigrilabris e de L. (L.) insularis coincidem com aquelas
previamente descritas como tpicas do gnero Leucochrysa e de seu subgnero Leucochrysa; (2) com base nas caracters-
ticas da genitlia ( e ), morfologia larval e biologia (por exemplo, deposio de ovos em cachos), as duas espcies so
distintas, mas muito proximamente relacionadas, e (3) L. (L.) nigrilabris e L. (L.) insularis compartilham vrias caracters-
ticas com Leucochrysa (L.) varia e espcies afins, as quais incluem polimorfismo de colorao no estgio adulto e aspectos
da morfologia larval semelhantes. Entretanto, suas genitlias ( e ) diferem significativamente das de L. (L.) varia e
espcies afins descritas, de forma que esses dois conjuntos de espcies so considerados como distintos.

Resumen

El presente estudio taxonmico incluye: (i) redescripcin de Leucochrysa (Leucochrysa) nigrilabris (Banks) ( y ), con
base en el espcimen tipo y nuevo material y (ii) imgenes del tipo, polimorfismo de coloracin de adultos y genitalia (

Accepted by A. Contreras-Ramos: 18 Oct. 2013; published: 18 Dec. 2013 101


Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
y ) de Leucochrysa (L.) insularis (Walker). Para ambas especies se presenta: (iii) descripciones de las larvas, (iv) infor-
maciones biolgicas y (v) registros geogrficos. Tales informaciones son usadas para comparar las dos especies entre s y
con otras especies de Leucochrysa (Leucochrysa) que supuestamente estn estrechamente relacionadas. A partir de las
comparaciones, se concluye que: primero, las caractersticas de las larvas de L. (L.) nigrilabris y de L. (L.) insularis coin-
ciden con aquellas previamente descritas como tpicas del gnero Leucochrysa y del subgnero Leucochrysa; segundo,
con base en las caractersticas de la genitalia ( y ), morfologa larval y biologa (por ejemplo, deposicin de huevos en
racimos) las dos especies son distintas, pero estn estrechamente relacionadas y tercero, L. (L.) nigrilabris y L. (L.) insu-
laris comparten varias caractersticas con Leucochrysa (L.) varia y especies relacionadas, las cuales incluyen polimorfis-
mo de coloracin en la fase adulta y aspectos de la morfologa larval. Sin embargo, las genitalias ( y ) difieren
significativamente de las de L. (L.) varia y las especies relacionadas descritas, de forma que los dos conjuntos de especies
son considerados distintos.

Introduction

Our study provides a taxonomic treatment of two species -- Leucochrysa (Leucochrysa) nigrilabris (Banks) and
Leucochrysa (Leucochrysa) insularis (Walker). It is part of a series dealing with the systematics of the
leucochrysine subgenus Leucochrysa (Leucochrysa) (Tauber 2004, Mantoanelli et al. 2006, Tauber et al. 2011,
2013).
Previously, L. (L.) nigrilabris was known from a single male specimen (the type, from Colombia), whose
original description (Banks 1914-1915) included only external features of the body and wings. Subsequently, the
species received no additional descriptive work. Given the large diversity of similarly colored Leucochrysa
(Leucochrysa) species, and given the variability in adult coloration and markings, current information is
insufficient for secure identifications.
In contrast, the adults of L. (L.) insularis, a species reported from eastern USA and the Caribbean region, have
had modern taxonomic treatment (Adams 1977, Tauber 2004) and identifications are possible. Also, the larvae
have received some attention and the unusual biology of this species has been described (Jones 1929, 1941), but
there are insufficient data to make larval identifications or meaningful comparisons with other Leucochrysa
species.
In the past, systematic relationships were proposed for these two Leucochrysa species, but the corroborating
evidence was meagre. Some authors noted that the external features of L. (L.) nigrilabris and L. (L.) insularis
adults resemble those of the South American Leucochrysa (L.) varia (Schneider) [= L. (L.) varia-like species]
(Banks 1914-1915; also see Tauber et al. 2013). In addition, the two species themselves were considered either
very closely related or synonymous (Tauber 2004).
To help clarify the systematics of the subgenus Leucochrysa (Leucochrysa), we redescribe the L. (L.)
nigrilabris adults (male & female) on the basis of the type and newer material from Venezuela and Brazil, and we
provide images of the L. (L.) insularis type, as well as comparative information on the species color
polymorphism and genitalia. Also, we describe the larvae of L. (L.) nigrilabris (all instars) and L. (L.) insularis
(first and third instars), and provide new biological notes on both species. Finally, to elucidate the systematic
relationships of the two species with each other and within the subgenus, we consider three questions: (1) Are the
larval features of L. (L.) nigrilabris and L. (L.) insularis consistent with those of other species in the subgenus
Leucochrysa (Leucochrysa)? (2) Do adult and larval features support the synonymy and/or proposed close
relationship between the two species? (3) Should the two species be aligned with the L. (L.) varia-like species?

Materials and methods

All terminology and methods are identical to those used previously (Mantoanelli et al. 2011, Tauber et al. 2013).
The specimens are from the following museums: American Museum of Natural History, New York, NY (AMNH);
The Natural History Museum (formerly British Museum of Natural History), London, England (BMNH); Bernice
P. Bishop Museum, Honolulu, HI (BPBM); Carnegie Museum of Natural History, Pittsburgh, PA (CMNH); C. P.
Gillette Museum of Arthropod Diversity, Colorado State University, Boulder, CO (COSU); Cornell University
Insect Collection, Ithaca, NY (CUIC); University of Guelph Insect Collection, Guelph, Ontario, Canada (DEBU);

102 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


Los Angeles County Museum, Los Angeles, CA (LACM); Museum of Comparative Zoology, Harvard University,
Cambridge, MA (MCZ); Museum of the Institute of Agricultural Zoology, Universidad Central de Venezuela,
Maracay, Aragua, Venezuela (MIZA); Royal Ontario Museum, Toronto, Ontario, Canada (ROM); Essig Museum,
University of California, Berkeley, CA (UCB); Bohart Entomological Museum, University of California, Davis,
CA (UCD); Museo Entomolgico Dr. Jos Manuel Osorio, Universidad Centroccidental Lisandro Alvarado,
Barquisimeto, Lara, Venezuela (UCOB); National Museum of Natural History (formerly United States National
Museum), Smithsonian Institution, Washington, D.C. (USNM).

Leucochrysa (Leucochrysa) nigrilabris (Banks, 1915)


Figs 112, 19

Allochrysa nigrilabris Banks, [1915] (Banks 19141915: 623) original description: [Colombia] St. Antonio, Colombia, 1,800
m., December (Fassl).
Leucochrysa nigrilabris (Banks). Banks (1944: 32) first reference to combination; Penny (1977: 23) species list.
Leucochrysa (Leucochrysa) nigrilabris (Banks). Brooks & Barnard (1990: 276) subgeneric designation, species list; Tauber
(2004: 1140) possible synonymy with Leucochrysa (Leucochrysa) insularis (Walker); Oswald (2007) catalog listing.

Type specimen. One type, from San Antonio, Colombia, MCZ (male, examined). We assume that the locality St.
Antonio refers to the city of San Antonio in the Department of Tolima. Banks (19141915) did not state how
many specimens he used to write his description; it is likely that he had only one. There is only one type in the
MCZ and none in the AMNH, USNM, or the Academy of Natural Sciences of Philadelphia, where Banks
occasionally deposited types. Thus, we consider the specimen in the MCZ to be the holotype, by monotypy. Its
labels read: (1) St Antonio / Colombia S. Am [Banks hand]; (2) 1800 m / Dec [Banks hand]; (3) Collection
/ N. Banks; (4) Type [red, Banks hand]; (5) Type / 12004; (6) Allochrysa / nigrilabris / type / Bks [white,
red border, Banks hand]; (7) Genitalia / prep P. Adams / 1982.
The specimen is in fairly good condition; the body (thorax and head) are somewhat flattened, the forewings
and one hindwing are missing, and the terminalia are in a vial with glycerine. We took images that are now placed
in the MCZ Type Database (http://insects.oeb.harvard.edu/MCZ/index.htm).
Known geographical distribution. Colombia: Department of Tolima. Venezuela: States of Aragua, Mrida,
Portuguesa, Yaracuy. Brazil: State of Rio de Janeiro.
Diagnosis (adult). Externally, L. (L.) nigrilabris adults are distinguished from many Leucochrysa
(Leucochrysa) species by a dark brown to black labrum, white frons, a single, small spot on the forewing (brown
suffusion on the membrane around the distal Psm-Pcu crossvein), and all veins mostly green except inner and outer
gradates dark brown to black. However, these features are shared with other Leucochrysa (Leucochrysa) species,
including some L. (L.) varia-like species and L. (L.) insularis. Thus, a reliable identification requires
examination of the genitalia. The males of L. (L.) nigrilabris are distinguished by: (i) dense microtholi on sternites
38; (ii) small genital structures [gonarcal span of 0.400.65 mm; cf., 0.670.84 mm in L. (L.) varia]; (ii) a narrow,
elongate mediuncus that is well separated from the gonarcal bridge, and terminates in a long, slender hook and a
pair of large, inflated flanges; and (iii) terminus of S8+9 with an attached pair of flat, sclerotized, plate-like
projections with toothed mesal margins. Female L. (L.) nigrilabris have a doughnut-shaped spermatheca with a
broad, bent velum, a deep, relatively narrow invagination, a relatively long, robust spermathecal duct, and
apparently no bursal duct; the subgenitale has a well-sclerotized ventral process that extends perpendicularly from
the distal margin of the base.
Redescription (adult). Head (Fig. 1): 1.71.9 mm wide (including eyes). Frons, clypeus white to cream-
colored, frons without red or pink suffusion, clypeus with or without tinge of pink; genae reddish brown; labrum
black throughout; maxillary, labial palpi yellowish to cream-colored. Vertex with central area raised, green mesally,
golden brown to reddish brown laterally; lateral, frontal areas cream-colored, with prominent, dark red to dark
brown, scalloped to V-shaped mark along anterior margin, extending anteriorly to or between mesal margins of
scapes, extending laterally to eyes; post-ocular area cream-colored, without spots. Antenna: scape cream-colored
ventrally, with reddish brown mark of various sizes on distal, dorsolateral surface; pedicel cream-colored basally,
with brown band distally; flagellum cream-colored, with brown bristles; mesal-ventral surface of basal one to
seven flagellomeres lightly tinged with brown (variable); dorsal antennal fossae cream-colored.

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 103


FIGURE 1. Leucochrysa (L.) nigrilabris external features. A, B. Head, frontal; C. Head, dorsal; D. Body, dorsal E, F. Head and
thorax, dorsal [note the variation in the coloration of the mesoprescutum and mesoscutellum]; G. Head, thorax, lateral; H. Head,
prothorax, lateral [A, D, State of Yaracuy, Venezuela, female; B, C, FH, State of Rio de Janeiro, Brazil, female; E, State of
Aragua, Venezuela, female].

Thorax (Figs 1DH): Cervical sclerite with large, red mark laterally. Pronotum 0.920.98 mm long, 1.411.44
mm wide, light green, with elongate, dark red stripe on anterolateral margin, diffuse, light brown, sublateral spots
anteriorly, posteriorly. Mesonotum, metanotum variable, with two distinct morphs [Figs 1D, E (brown), 1F
(white)]. Brown morph: mesonotum, metanotum dark brown throughout, except anterior margin of prescutum
cream-colored, with pair of sublateral, dark brown spots; mesoscutellum slightly lighter brown than remainder of

104 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


notum; metascutellum black throughout. White morph: mesonotum mottled black to brownish black, anterior
margin of prescutum cream-colored, with pair of sublateral, dark brown spots, posterior margin with cream-colored
areas, mesoscutellum entirely white; metanotum largely dark brown to black, metascutum with large, white
posteromesal patch; metascutellum black throughout.

FIGURE 2. Leucochrysa (L.) nigrilabris wings. A, B. Forewing, two specimens to illustrate relatively minor variation; C.
Hindwing [A, State of Yaracuy, Venezuela; B, C, State of Aragua, Venezuela, all female]. Scale applies to all three images.

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 105


FIGURE 3. Leucochrysa (L.) nigrilabris male abdomen, mature specimen. A. Second and third abdominal segments, lateral
[note absence of microtholi from S2, dense microtholi on S3]; B. Terminal segments, lateral; C. Terminal segments, dorsal
[note large brown spots on T5, T6, base of T7 (arrow)]; D. Terminal segments, dorsal [note dorsally fused T9+ectoproct,
without suture or distal cleft. The U-shaped darkening is the distal margin of S8+9 below]; E. Callus cerci; F. Fused sternite
8+9, ventral, with sclerotized terminal plates extended distally [all, Aragua, Venezuela]. Abbreviations: c.c., callus cerci; d.a.,
dorsal apodeme of ninth tergite+ectoproct; s.p., sclerotized plate at tip of S8+9; S2, S3, second and third sternites; S8+9, fused
eighth and ninth sternites (in parentheses, out of focus through dorsal surface); tr, trichobothria; T5, T7, T8, fifth to eighth
tergites; T9+ect, fused ninth tergite and ectoproct.

106 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


FIGURE 4. Leucochrysa (L.) nigrilabris male genitalia, mature specimen. A. Gonarcus, posterodorsal (mediuncus removed);
B. Gonarcal complex, posterior (view into gonosaccus from behind, through membrane) [note the mediuncal beak and flanges
extend forward (away) and downward]; C. Gonarcal complex, dorsal [view into gonosaccus from above, unobscured by
membrane, base of mediuncus in focus, mediuncal beak present (extending forward), but out of focus]; D. Gonarcal complex,
frontal [note the gonosaccus is below the mediuncal flanges; the gonosetae are within the gonosaccus]; E. Gonarcal complex,
lateral; F. Hypandrium internum (comes very reduced or absent) [all, Aragua, Venezuela]. Abbreviations: bk, beak-like tip of
mediuncus; fl, mediuncal flange; gc, gonocornu, encased in membrane; g.a., gonarcal apodeme; g.b., gonarcal bridge; g.p.,
gonarcal process (membrane attachment); gsac, gonosaccus; gse, gonosetae (within gonosaccus); mu, mediuncus (fused rods
and beak); mu-b, sclerotized base of mediuncus.

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 107


FIGURE 5. Leucochrysa (L.) nigrilabris male abdomen, teneral specimen. A. Terminal segments, lateral (arrows mark the
large brown spots on T6, base of T7); B. Gonarcal complex, frontal; C. Gonarcal complex, lateral [note thorny membrane over
gonarcal bridge and gonocornua]; D. Gonarcal complex, lateral (thorny membrane above gonarcus partially removed); E. Tip
of sternite 8+9, dorsal, with sclerotized terminal plates extended distally, hypandrium internum (attached to small membrane
that had been everted beneath the sclerotized plate); F. Sclerotized plate at tip of S8+9, lateral (all, Aragua, Venezuela).
Abbreviations: bk, beak-like tip of mediuncus; c, comes; cleft, cleft separating distal margins of T9+ect; c.c., callus cerci; d.a.,
dorsal apodeme of T9+ect; fl, mediuncal flange; gc, gonocornu; gs, gonarcus; gsac, gonosaccus with robust gonosetae on
bulbous chalazae; g.a., gonarcal apodeme; g.b., gonarcal bridge; h.i., hypandrium internum; mu-b, sclerotized base of
mediuncus; mu-d, elongate distal section of mediuncus; s.p., sclerotized plate at tip of S8+9; S8+9, fused eighth and ninth
sternites; S9, ninth sternite (tip); T7, seventh tergite; T9+ect, fused ninth tergite and ectoproct..

108 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


FIGURE 6. Leucochrysa (L.) nigrilabris female abdomen. A. Fifth to terminal segments, lateral; B-D. Seventh to terminal
segment, lateral [A, B, mature specimens, State of Rio de Janeiro, Brazil; C, D, teneral specimens, State of Aragua, Venezuela].
Abbreviations: g.l., gonapophysis lateralis; sg, subgenitale; sp, spermatheca; S7, seventh sternite; T5, T8, fifth and eighth
tergites; T9+ect, fused ninth tergite and ectoproct.

Wings (Fig. 2) Forewing 18.819.9 mm long, 6.56.9 mm wide (at widest point); ratio of length: maximum
width = 2.62.7:1. Costal area moderately broad; tallest costal cell (#910) 1.61.7 mm tall, 3.03.1 times width,
0.20.3 times width of wing (midwing). First intramedian cell quadrangular, width (anterior margin) 1.41.7 times
width (anterior margin) of third median cell, 0.91.1 times length of posterior margin of m3; length of basal vein (=
ma, median arculus) 2.16.2 times greater than length of distal vein. First radial crossvein distal to origin of radial
sector (Rs); radial area (between Radius and Rs) with single row of 1718 closed cells; tallest cell (#8) 2.32.4
times taller than wide. No crassate veins; 56 b cells (= cells beneath Rs, not including an inner gradate vein). Two
series of gradate veins; 1012 inner gradates, 810 outer gradates. Height of fourth gradate cell 4.35.3 times
width. Eight to nine b' cells (cells beneath pseudomedia after second intramedian cell). Three intracubital cells (two
closed). Membrane clear except stigma opaque, with dark brown mark basally, distal two crossveins of last b cell
(distal Psm-Pcu crossveins) surrounded by dark brown suffusion, and (some specimens) small amount of brown
suffusion basally. Veins green, except distal two crossveins of last b cell, inner and outer gradates dark brown to
black, sometimes anterior tips of basal R-Rs crossveins, base of Rs, basal Rs-m, basal medial crossveins, tips of
posterobasal marginal veins lightly darkened.
Hindwing 16.717.7 mm long, 5.55.8 mm wide. Two series of gradate veins; 910 inner, 89 outer; 1516
radial cells (counted from origin of Radius, not false origin). Five to six b cells (including small b1 cell); seven to
nine b' cells beyond second intramedian cell; two intracubital cells (one closed). Membrane clear; stigma with
pronounced brown mark basally. Veins light green.

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 109


Abdomen (Figs 38): Tergites with mostly short, slender setae throughout, sternites with longer, slender setae;
microsetae dense; pleural region with setae small, very sparse, microsetae very small. Tergites narrow, roughly
rectangular, with rounded or irregular margins. Spiracles oval externally; atria not enlarged. Sternites S2S5 longer
than wide (lateral view). Coloration: mostly green, with diffuse yellow dorsal stripe; tergites T5, T6, base of T7
each with large dark brown to black spot, bordered with yellow and patches of red; callus cerci white; setae,
trichobothria golden.

FIGURE 7. Leucochrysa (L.) nigrilabris female abdominal and genitalic structures. A. Callus cerci; B. Subgenitale, posterior;
C. Spermatheca, lateral; D. Genitalia, ventral [A, B, D, State of Rio de Janeiro, Brazil, mature specimen; C, State of Aragua,
Venezuela]. Abbreviations: base, sclerotized base of subgenitale; b.c., bursa copulatrix; b.gl., bursal gland; inv, spermathecal
invagination; lobe, rounded lobe of subgenitale; sg/b.c., heavy folded membrane [distally (right): dorsum of subgenitale,
proximally (left): venter of bursa copulatrix]; sp, spermatheca; sp.d., spermathecal duct; tr, trichobothria; vel, velum.

Male. S6, S7 height ca 1.11.2 times length (lateral view). Microtholi dense on S3S8, absent from S1S2, S9.
Callus cerci oval (ca 1.31.5x taller than wide), greatest diameter 0.220.26 mm, with 2635 trichobothria of
variable length. T9+ectoproct soft, lightly sclerotized, rounded posterodorsally, truncate distally, broadly fused
mesally, midline without distal cleft (mature specimen; cleft present in teneral specimen); ventral section tapering
proximally, extending beyond midline of S9; dorsal apodeme lightly to moderately sclerotized, straight, extending
ca half distance along dorsal margin of ectoproct; small, lightly sclerotized apodeme around proximal margin of
callus cerci. S8+9 fused, without suture, with clear intersegmental demarcation throughout; dorsal margin with
lightly sclerotized, straight apodeme; S8 1.7x taller than long, ca one-half (0.470.51x) length of S8+9; S8+9
(lateral view) with proximal margin relatively straight, dorsal surface straight, sloping ventrally to distal margin;
terminus rounded; tip with pair of flat, sclerotized, plate-like projections with mesal edge toothed, lateral edge
rounded. Gonarcus arcuate; bridge straight, narrow mesally (0.19 mm), curved abruptly at exterior margin of
gonocornua; gonarcal bridge, gonocornua covered with robust membrane bearing dense gonocristae; gonocornua
short (length, 0.07 mm), extending forward from edge of gonarcal bridge, stout basally, tapering to rounded apex;

110 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


distance between inner bases of gonocornua 0.07 mm basally, distance between tips 0.12 mm; gonarcal lateral
apodeme long, relatively narrow (0.60 mm long, 0.20 mm wide), not extending above bridge, curved distally
(lateral view); distance between apodemes distally 0.72 mm. Gonarcus, between lateral apodeme and gonocornu,
with moderately sclerotized, elongate posteroventral projection holding membrane that extends beneath
mediuncus. Mediuncus well separated from gonarcus, with basal membranous connection stout; basal section of
mediuncus consisting of heavily sclerotized platform with distal projections laterally, distal section consisting of
well sclerotized, ventrally-projecting, smooth plate terminating in sharp, curved, elongate, mesal beak, pair of large
lateral lobes; heavy membrane from gonarcus extending beneath mediuncus, recurving above beak to form
gonosaccus with pair of elongate, lateral fields of robust gonosetae on bulbous chalazae; fields of gonosetae on
surface of membrane facing mediuncus; each field with ca 1520 gonosetae. Hypandrium internum broadly V-
shaped, with comes not visible (mature specimen); arm 0.200.22 mm long, distal span between arms 0.19 mm.

FIGURE 8. Leucochrysa (L.) nigrilabris colleterial complex. A. Colleterial complex, lateral; B. Base of colleterial complex,
lateral (pulled posteriorly); C. Transverse sclerite, posterior. [all, State of Rio de Janeiro, Brazil, mature specimen].
Abbreviations: acc. gl., colleterial accessory gland; acc.gl.d., duct of colleterial accessory gland; c.d., common duct (colleterial
reservoir and accessory gland to transverse sclerite); col.gl., colleterial gland; col.r., colleterial reservoir; tr.s., transverse
sclerite.

Female. S6 height ca 0.670.92 times length, S7 height ca 0.550.65 times length. Callus cerci round,
diameter ca 0.170.19 mm, with 2934 trichobothria. T8 roughly elliptical (lateral view) with rounded corners,
similar in depth to T6. T9+ectoproct elongate, vertically erect; ventral margin slightly rounded, extending below
level of gonapophyses laterales. S7 with dorsal margin straight, distal end tapering abruptly; terminus truncate,

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 111


unmodified, with terminal (posteroventral) setae slightly more numerous, longer than in other areas. Gonapophysis
lateralis rounded throughout, ~0.480.52 height of T9+ectoproct; inner membranous surface slightly expandable,
with vertical patch of small, delicate setae arising from slightly swollen membrane. Colleterial complex consisting
of membranous gland, connected to colleterial reservoir via broad duct, and large accessory gland, both opening to
exterior via narrow common duct above transverse sclerite; colleterial gland elongate, delicate; colleterial reservoir
shorter, robust; accessory gland probably bulbous (torn in our specimens), elongate, granular, with multiple forks
distally; transverse sclerite broad, flat, bent distally, with relatively dense, elongate, hair-like setae. Spermatheca
simple, doughnut-shaped, with broad, flat, L-shaped velum, maximum diameter 0.260.31 mm, total length
(spermatheca + velum) ca 0.35 mm; spermathecal invagination deep, narrow, with rounded terminus; velum with
elongate slit opening to bursa copulatrix; tip of velum bending slightly into bursa copulatrix; bursal duct very small
or absent; spermathecal duct relatively long, ca 1.131.51 mm, originating from side of spermatheca distal to slit in
velum, basal half well sclerotized, tapering throughout, with lateral bend, then U-shaped loop; distal ca half
narrow, brushy throughout, with curve, followed by distinct swelling mesally, with densely hirsute, swollen
terminus. Bursa copulatrix saccular, with heavily textured surface distally (below subgenitale), becoming smoother
anteriorly, with pair of elongate, tubular bursal glands attached to distal end, connections appearing unremarkable.
Subgenitale with sclerotized surface smooth (unfolded), attached basally to heavy, transversely folded basal
membrane (base of subgenitale then top of bursa copulatrix), with two projections, bilobed one on dorsodistal
margin, unilobed one on anteroventral margin base withdrawn above tip of S7 [with pressure, the withdrawn lobe
can be extruded].
Intraspecific variation (adult). Leucochrysa (L.) nigrilabris expresses considerable variation in head and
mesothoracic coloration, as well as wing size, shape and degree of brown suffusion surrounding various veins. The
most striking aspect of the variation is the polymorphism in mesothoracic coloration (Figs 1DF). Among the
specimens we examined, eight (4, 4) had brown mesoscutella, and five (2, 3) had white; both color forms
occurred among the Venezuelan specimens.
The male genitalia show considerable developmental variation (Compare Fig. 5 with Figs 3, 4). In the teneral
male, the gonarcus is weakly sclerotized and narrow; the mediuncal bridge appears wide (in lateral view) and is
tightly encased by the gonocristae-bearing membrane; the mediuncus is very narrow; and the hypandrium internum
is unexpanded [length of arms, 0.20 mmas in mature specimens; distal span between arms, 0.06 mmca 3x
narrower than mature specimens]. Females, too, show considerable variation with age and maturation. For
example, in the teneral female, the subgenitale is small, weakly sclerotized, and withdrawn, and the spermatheca is
shifted anteriorly from the distal wall; whereas in mature specimens, the subgenitale is large, robust, heavily
sclerotized and expanded, so that the spermatheca appears surrounded by the distal wall (Figs 6, 7B).
Description (First Instar, Semaphoront A). Body (Fig. 10). 2.83.0 mm long. All setae smooth.
Head (Figs 10AC). 0.550.56 mm wide, 0.380.39 mm long; mandibles 0.590.61 mm long; ratio, mandible
length : head width, ~1.09:1. Epicranial marking brown, paired, extending along entire basal margin of cranium,
laterally extending to eye, mesally becoming confluent with base of postfrontal marking. Postfrontal marking
slightly darker brown than epicranial marking, elongate, extending to base of antenna. Frontal marking V-shaped,
arms extending forward through dark tentorial pits becoming confluent with intermandibular marking.
Intermandibular marking broken mesally, extending laterally from tentorial pits to base of mandibles, anteriorly to
tip of clypeus. Gena with single, elongate, brown stripe extending from base of eye to posterior edge of cranium.
Labial palpus cream-colored basally, light brown distally. Mandibles amber to brown. Antenna light brown to
cream-colored. Venter cream-colored to white mesally, light brown laterally, with brown transverse stripe distally;
cardo, stipes brown.
All cephalic setae straight, with pointed terminus. Primary setae S1S12 present; S1, S11 long; S2, S3, S5
short. Labial palpus: basal segment with two setaeone lateral, one ventral; midsegment with two distal setae
one lateral, one mesal. Mandible with one basolateral seta; maxillae with basal seta. Palpiger with two basal setae.
Scape without setae; pedicel with one small distal seta; flagellar seta slightly longer than one-half length of
segment.
Thorax (Figs 10D, G, H). Notum with diffuse, reddish brown, submesal markings; Setae (LS) on lateral
tubercles (LTs) dark brown; dorsal setae light brown. Legs mostly light brown to brown; trochanters, tarsi white to
cream-colored; claws, empodia brown. LS gently curved throughout, with tips pointed, fine; all other setae straight
to gently curved, with terminus pointed.

112 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


T1: LT with two long LS, without microseta between. First primary sclerite (Sc1) light brown, large, with
associated seta (S1Sc1) long, arising from small chalaza; S2Sc1 short, anterior to S1Sc1. S1 long, arising from
small chalaza; S2 absent; S3, S4, S5 short.
T2: Spiracles prominent, tubercle-shaped, apex sclerotized, dome-shaped, amber-colored. LT with three long
LS. Sc1, Sc2 small; associated setae (S1Sc1, S2Sc1 and S1Sc2) very small; Sc3 intermediate-sized, light brown,
with associated seta (S1Sc3) small. Posterior subsegment with pair of short setae mesal to Sc3, row of four setae
posterior to Sc3 (mesal pair short, lateral pair long, arising from robust chalazae).

FIGURE 9. Prothorax of third instar of four Leucochrysa (Leucochrysa) species. A. L. (L.) nigrilabris; B. L. (L.) insularis; C.
L. (L.) varia; D. L. (L.) boxi [A, C, D, State of Rio de Janeiro, Brazil; B, North Carolina, United States].

FIGURE 10. Leucochrysa (L.) nigrilabris first instar. A. Head, dorsal; B. Head, ventral; C. Head, lateral; D. Thorax, dorsal; E.
Abdominal segments 1 to 5, dorsal; F. Abdominal segments 4-10, dorsal; G. Body, lateral; H. Body, ventral [all, State of Rio de
Janeiro, Brazil]. Abbreviations: Ax, abdominal segment number; AxRx, setal row number.

T3: LT with three long LS. Sc1 small, with S1Sc1 very small; Sc2 light brown, with S1Sc2 small. Anterior
region with pair of short setae; posterior region with four long setae arising from robust, fleshy chalazae.
Abdomen (Figs 10EH). White to cream-colored; dorsum of A1A3 with diffuse, submesal, reddish brown
band; dorsum of A4, A6A8 each with pair of reddish brown submesal spots. LTs each with two long LS (A2A7),
without microseta between; LS straight, with pointed tip, dark brown; submedian setae (SMS), setae (LDS) on
laterodorsal tubercles (A1A5) hooked, light brown. A1, A6A8: Spiracle without associated seta (SSp); A2A5:
each spiracle with small to minute SSp. A1A5: Laterodorsal tubercles (LDTs) each with two long, hooked LDS,
one microseta between LDS. A1: Single row of four SMS between LDTs. A2A4: Anterior row of four SMS,
posterior row of two SMS, both between LDTs. A5: Anterior row of two to three SMS, posterior row of two SMS,
both between LDTs. A6: Two short, straight setae anteriorly, two intermediate-length, hooked setae posteriorly;
LDT with one long, hooked LDS, one short, straight LDS. A7: LDT with one intermediate-length, straight LDS,
two very short LDS; dorsum with pair of minute setae anteriorly. A8: LT with one short, straight LS, two to three
small setae, base of LT with spiracular tubercle projecting from anterior surface; spiracular base and sclerotized
ring elongate, projecting posteriorly; dorsum with three to four short, straight setae between LTs. A9: Dorsum with

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 113


three rows of short, straight setae (anterior two minute, middle four short, posterior four slightly longer). A10:
Apparently without setae; terminus light brown, with small eversible pouch.
Description (Second & Third Instars, Semaphoront B). Body (Figs 9, 1112). L2: 4.74.9 mm; L3: 8.69.0
mm long. Dorsum cream-colored, with dark brown markings.

FIGURE 11. Leucochrysa (L.) nigrilabris second instar. A. Head, dorsal; B. Head, lateral; C. Thorax, dorsal; D. Abdominal
segments 1 to 4, dorsal; E. Abdominal segments 410, dorsal; F. Body, lateral [all, State of Rio de Janeiro, Brazil].
Abbreviations: Ax, abdominal segment number; AxRx, setal row number.

Head (Figs 11A, B, 12A, B, G, H). L2: 0.810.83 mm wide, 0.530.58 mm long; mandibles 0.930.96 mm
long; ratio mandible length : head width 1.141.17:1. L3: 1.31.4 mm wide, 0.860.91 mm long, mandibles 1.5
1.7 mm long; ratio mandible length : head width, 1.191.23:1. Epicranial marking paired, brown to dark brown,
covering entire basal portion of cranium, laterally becoming lighter brown, extending to eye, mesally becoming
confluent with base of postfrontal marking. Postfrontal marking slightly darker brown than epicranial marking,
elongate, extending to base of antenna. Frontal marking V-shaped, arms extending forward through dark tentorial
pits, then curving to base of mandibles becoming confluent with intermandibular marking. Intermandibular
marking brown, divided into three sectionstwo lateral sections at mesal base of mandibles, mesal section
between, but separated from two lateral sections; clypeolabral region partially covered by intermandibular
marking. Gena with single, elongate, brown stripe extending from base of eye to posterior edge of cranium; region
around eyes cream-colored; stemmata, except posteroventral margin, surrounded by brown. Labial palpus: basal
segment light brown; mesal segment light brown basally, darker distally, with well defined annulations; terminal
segment dark brown. Mandibles brown basally to dark brown at tip. Antenna light brown to brown. Ventrolateral
cranial surface light brown. Cardo with light brown mark basally, dark brown mark distally, narrow; stipes light
brown to tan, narrow, elongate. Mentum white to cream-colored, with tinge of light brown anteriorly; palpiger light
brown. Cervix cream-colored, with prominent lateral patches of brown.
Anterior margin of head with slightly rough, obtuse lateral edges; posterior section of cranium spiculate. All
cephalic setae smooth, straight, with acute tip; S1S12 present; S1, S11 long; S2, S3, S5 short. Labial palpus: basal
segment with three setae, midsegment with distal annulation bearing four setae, remaining annulations with ca six
(L2) or seven (L3). Mandible with one basolateral seta; maxillae with basal seta. Palpiger with two basal setae.
Scape with one distolateral seta; pedicel with one small distal seta; flagellar seta ca one-half length of segment.
Thorax (Figs 9, 11C, F, 12C, H, I). LTs white to cream-colored, with diffuse to very pale, reddish brown
markings at dorsal base submesally; LS mostly dark brown; large sclerites Sc1 (prothorax), Sc3 (mesothorax), Sc2
(metathorax) light amber, shiny. Venter white to cream-colored, with small, diffuse, rosy, submesal patches on each
segment. Spiracles amber to light amber; setae mostly brown or light brown. Legs: coxa with dark brown marking

114 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


on dorsal connection to base, pair of brown marks, one anterior, one posterior; trochanter with pair of dark brown
marks, one anterior, one posterior; femur brown dorsobasally, light brown distally; tibia brown throughout; tarsus
light brown; claws curved, large, dark brown; empodia dark brown.
LTs smooth; LS smooth, except very fine tips sometimes slightly salebrose, seldom sparsely and finely barbed,
slender, gently curved throughout, tips tapering to very fine terminus; tips of distal LS mostly pointed, tips of
lateral LS mostly with small hook. All dorsal, ventral setae smooth.

FIGURE 12. Leucochrysa (L.) nigrilabris third instar. A. Head, dorsal; B. Head, ventral; C. Thorax, dorsal; D. Abdominal
segments 1 to 3, dorsal; E. Abdominal segments 37, dorsal; F. Abdominal segments 6-10, dorsal; G. Head, lateral; H. Body,
ventral; I. Body, lateral [all, State of Rio de Janeiro, Brazil]. Abbreviations: Ax, abdominal segment number; AxRx, setal row
number.

T1: All dorsal, ventral setae straight; Sc1 large, extending laterally to dorsal surface of LT, with S1Sc1 long,
slender, S2Sc1 very short, slender. Sc2 transparent, oval-shaped, ca one-third length of Sc1. LT (L2) with 1921
LS (ca 10 apical, ca 911 lateral); LT (L3) with ca 2730 LS (ca 10 apical, ca 1720 lateral). Primary setae S1
medium-length, S2, S3, S4, S5 short, slender; no secondary setae.
T2: Anterior subsegment without secondary setae between spiracles; anterior sclerite (Sc1) with three
associated setae (S1Sc1, S2Sc1, S3Sc1); spiracles prominent, with opening sclerotized, dome-shaped, amber-
colored ring on small tubercle; walls of chamber parallel, not bulbous; SSp absent. Posterior subsegment with
anterior sclerite (Sc2) bearing two very small associated setae (S1Sc2, S2Sc2); Sc3 with small associated seta
(S1Sc3); anterior row with two intermediate-length, slender, straight setae; posterior row (L2) with ca 56 setae
(34 long, robust, 12 slender, hooked or straight), arising from large, plump chalazae; posterior row (L3) with ca
1114 setae (68 long, robust, 57 slender, hooked or straight), arising from large, plump chalazae. LT (L2)
bearing ca 1719 LS (ca 89 apical, ca 910 lateral, mostly straight); LT (L3) bearing ca 26 LS (ca 10 apical, ca
1517 lateral).

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 115


T3: Sclerites (Sc1, Sc2) each with one associated setae (S1Sc1, S1Sc2); sometimes small seta on anterior fold,
lateral to Sc1. Anterior row with one to two short (L2) or two to four intermediate-length (L3), slender, straight
setae; posterior row (L2) with ca 12 long, robust, straight or hooked setae arising from large, plump chalazae;
posterior row (L3) with ca 1516 setae (ca ~1214 long, robust, 24 intermediate-length, slender, straight or
hooked), arising from large, plump chalazae. LT (L2) bearing ca 1920 LS (ca 9 apical, 1011 lateral, mostly
straight). LT (L3) bearing ca 2125 LS (ca 9 apical, 1216 lateral).
Abdomen (Figs 11DF, 12DF, H, I). Dorsum, venter white to cream-colored; anterior segments without dorsal
markings; A6A8 with diffuse, reddish dorsal marks; A10 with small, brownish mesal spot distally. Venter of A2
A4 with diffuse, light reddish sublateral patches; A10 with pair of brown, submesal spots. LS on A2A3 light
brown, A4A7 dark brown; SMS light brown to dark amber.
All setae smooth. SMS of A1A6 hooked, slender, arising from plump chalazae; SMS longest, chalazae largest
on A1, A2, becoming smaller posteriorly. A1A5: LDT plump, with two long, slender, hooked LDS, one microseta
between LDS. A1A8: Spiracle small, slightly raised, dome-like, with minute to very small associated seta (SSp?)
anteriorly or mesally.
A1: Single transverse row of eight (L2) or ca nine to ten (L3) SMS mesal to LDTs.
A2A3: LT broad, plump, fleshy, with seven (L2) or eight (L3) long, robust LS arranged transversely along
dorsal surface, ca two (L2) or four (L3) shorter, straight setae basally. L2: Dorsum with anterior row of four SMS,
mesal row of four, posterior row of four (A2) or six (A3) between LDTs. L3: Dorsum with anterior row of ca ten
(A2) or ~12 (A3) SMS, mesal row of four (A2, A3), posterior row of ca 12 (A2, A3) between LDTs.
A4A5: LT tuberculiform, with ca five (L2) or 12 (A4, L3), 910 (A5, L3) robust, hooked or straight LS
apically, dorsally, ca four (L2, L3) short, straight LS basally. LDS intermediate length, hooked. L2: Dorsum with
anterior row of four (L2) intermediate-length SMS, mesal row of four, posterior row of four between LDTs. L3:
Dorsum with anterior row of ca 12 (A4) or ca nine (A5) intermediate-length SMS, mesal row of four, posterior row
of ca ten between LDTs.
A6: LT tuberculiform, with ca five (L2) or six (L3) robust, straight LS apically, ca three short, straight LS
basally. LDT with one long, hooked LDS, two to three short, straight LDS. Anterior row of three (L2) or five (L3)
intermediate-length SMS, no other SMS.
A7: LT tuberculiform with posterior projection, with ca five (L2) or six (L3) robust, straight LS apically, ca
three short, straight LS basally. LDT with one intermediate-length, straight LDS, one to two short, straight LDS.
Two pairs of very small setae between spiracles.
A8: LT with four to five short, straight LS (two short, two to three very short). Row of four to five very short
setae between spiracles. Posterior margin with ca four (L2) or seven to eight (L3) very short setae.
A9: Two transverse rows of ca four to six short setae.
A10: Dorsum with ca four to five very short setae.
Intraspecific variation (larva). Three variable features warrant notation. (i) As with the larvae of all chrysopids
we have studied, L. (L.) nigrilabris larvae exhibit considerable bilateral asymmetry in the numbers and size of setae
across the dorsum of body. Odd numbers of setae are common. (ii) The left side of the first abdominal segment (A1)
of one L3 specimen has a fleshy, lateral protuberance (LT?) bearing an elongate, robust, hooked seta. This feature is
unusual because LTs rarely occur on the A1 (e.g., in some Chrysopa spp., see Principi 1940, Tsukaguchi 1978), and
they have not been reported for leucochrysine larvae. Moreover, LTs almost always bear more than one elongate
seta. Thus, we do not know whether this structure is homologous to those on the larvas subsequent segments (A2
A7) and/or to those on the Chrysopa A1. (iii) Among the first instars, the elongate setae (LS) on the apex of the
lateral tubercles are mostly very long, straight, and tapering to pointed tips; occasionally some are hooked.
Biological notes. Adults were collected in disturbed habitats (e.g., coffee plantation, roadside, along trail) at
the edges of mature forests. One female specimen, collected in the State of Rio de Janeiro, was maintained in the
laboratory for ca two weeks. During that time she laid two batches of eggs. Each batch consisted of a hanging
cluster of ca 1520 eggs attached to the ends of a series of long intertwined stalks.
The L. (L.) nigrilabris eggs were maintained at room temperature; hatching occurred within ca ten days. The
larvae applied and carried dense packets of trash [in this case, frozen & thawed eggs of Ephestia kuehniella
(Zeller)] on their dorsa. The first moult occurred ca six days after hatching, and the second moult ca seven days
after the first; cocoons were spun ca 10 days later. Four pharate adults emerged from their cocoons; only one
completed ecdysis to the adult stage.

116 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


Specimens examined (adults and larvae, in addition to the type listed above). VENEZUELA. Aragua:
Parque Nacional Henri Pittier (formerly Parque Nacional Rancho Grande), 1100 m, 1113.ii.1969, J. Salcedo (1,
MIZA); idem, 22.v.1967, L. Rodriguez V. (1, MIZA, determined by Adams 1979); idem, 9.xi.1990, V. Savini,
trampa de luz (1, MIZA); idem, 5.vi.1958, F. Fernndez Y. (MIZA, determined by Adams 1979]; idem, 16
31.x.1966, S. S. & W. D. Duckworth (5, USNM); idem, 1021.x.1969, Duckworth & Dietz (1, 1, USNM);
idem, 8.vi.1967, R. W. Poole (2, CUIC); idem, 5.viii.1967, R. W. Poole (1, CUIC); idem, 20.ix.1973, B.
Villegas (1, UCD); idem, 2425.i.1996, white light, J. &. A. Skevington (2, 2, DEBU); idem, 1150 m, at
lights on bldg, 24.ii12.iii.1996, S. Marshall (1, 1, DEBU); El Limn, 450 m, 20.iv.1964, E. Osuna (MIZA,
determined by Adams 1979); Mrida: Mrida, el. ca 4500 ft., 8.5942N / 71.1429W, 2223.ii.1994, J. P. Donahue
(1, LACM); Portuguesa: Crdoba 800 m, 2429.xi.2008, R. Pz, Malaise Trap (1, UCOB); Yaracuy: La
Cumbre, Chivacoa Council, 1075 m, 1014N / 6858W, 22.vii.2012, in Coffea arabica, F. Sosa, manual net (1,
UCOB); idem, 1200 m, 1016N / 6856W, 11.ii.2013, in Coffea arabica, F. Sosa, manual net (1, UCOB); idem,
18.iv.2013, in Coffea arabica, F. Sosa, C. Martins and F. Daz, manual net (2, 1, UCOB); El Candelo, Cocorote
Council, 1309 m, 1020N / 6849W, 18.iv.2013, in forest, F. Sosa, C. Martins and F. Daz, manual net (1,
UCOB). BRAZIL. Rio de Janeiro: Sta. M. Madalena, Terras Frias, 28.x.2003, G. S. Albuquerque, M. J. Tauber, C.
A. Tauber Expedition, Oct.Nov. 2003 [1, 3?sex (lab-reared, pharate adults, incompletely emerged), 9 larvae
(Tauber Lot 2003:067, UCB); 15 larvae (Albuquerque Lot 2003:021, UENF)].

Leucochrysa (Leucochrysa) insularis (Walker, 1853)


Figs 9, 1322

L. (L.) insularis taxonomic history & geographic distribution. Adams (1977) and Tauber (2004) provided the
taxonomic history of the species and redescribed the adults based on the type specimen and other material. The
type of L. (L.) insularis (male, BMNH) is in relatively poor condition; it appears to be teneral, and its body was
damaged by dermestids. See images (Fig. 13).
Adams (1977) and Tauber (2004) also illustrated the male and female genitalia and documented significant
geographic variation in the species adult coloration. To help facilitate comparisons, here we provide images of the
L. (L.) insularis adult color polymorphism (Fig. 14), wing markings (Fig. 15), and genitalia (: Fig. 16; : Figs 17,
18).
Leucochrysa (L.) insularis occurs throughout the eastern and central states of the United States (north to
Massachusetts and Nebraska, south to Texas and Florida), eastern Mexico [Hidalgo, San Luis Potos, Tamaulipas,
Veracruz (new state record)], and the West Indies (Cuba, Dominican Republic and Jamaica). Thus, L. (L.) insularis
and L. (L.) nigrilabris appear to have adjacent, allopatric distributions.
Jones (1929) provided a well-illustrated larval description of L. (L.) insularis [as Nodita virginica (Fitch)].
Although it does not specify which instar (2nd or 3rd) is described, it contains fascinating information on both the
morphology and biology of the larvae. The descriptions below are based on new material. The first instars were
teneral and somewhat shriveled; their structures were not measured; many small setae were not visible. The third
instar was teneral and not measured. For both instars, characteristics are as described above for L. (L.) nigrilabris,
except where noted.
Description (First Instar, Semaphoront A).
Head. 0.54 mm wide, 0.40 mm long; mandibles 0.56 mm long; ratio, mandible length : head width, ~1.02:1.
Thorax. T1: S1 long, arising from large chalaza; S3, S4, S5 very short.
Abdomen. A6: Two short, straight setae anteriorly, no setae posteriorly.
Description (Third Instar, Semaphoront B).
Body. Dorsum cream-colored to brown, with brown sclerites.
Head (Figs 20AD). 1.32 mm wide, 0.76 mm long, mandibles 1.481.53 mm (right-left) long; ratio mandible
length : head width, 1.12:1.0. Genal marking light brown, diffuse, extending broadly toward eye and ventral margin
of cranium. All stemmata, including posteroventral one, surrounded by brown outline. Posterior section of cranium
densely spiculate.
Thorax (Figs 20D, E). Large sclerites Sc1 (prothorax), Sc3 (mesothorax), Sc2 (metathorax) brown, shiny;
markings on venter not visible. Setae associated with small sclerites between segments and subsegments not

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 117


readily observed. LTs spiculate; LS smooth basally becoming slightly salebrose distally, with tips mostly curly,
sometimes with small hook.
T1: Sc1 large, extending laterally to base of LT; S1Sc1 long, robust, S2Sc1 very short. LT with ca 22 LS (ca 10
apical, ca 12 lateral). Primary setae S1 long, robust; S2 short; S3, S4, S5 very short. T2: Posterior row with 6 setae
4 long, robust, straight to curvey, arising from 2 pairs of large, plump, juxtaposed chalazae, and 2 short, straight,
arising from mesal margin of mesal chalazae. LT bearing 2024 LS (8 apical, 1416 lateral). T3: Anterior row with
two intermediate-length, robust setae; posterior row with 12 long, robust, hooked setae arising from large, plump
chalazae. LT bearing ca 2023 LS (89 apical, 1115 lateral). [Note: The ranges in setal counts are derived from
paired tubercles.]

FIGURE 13. Leucochrysa (L.) insularis holotype (male), Jamaica, BMNH. A. Head, prothorax, dorsal; B. Head, frontal; C.
Head, prothorax, lateral; D. Forewing; E. Hindwing; F. Terminal segments of abdomen, lateral; G. Terminus of ninth abdominal
sternite, ventral; H. Labels.

Abdomen (Figs 20E, F). SMS of A1A6 robust, hooked, arising from plump chalazae; A1A5: LDT plump,
with two long, hooked, robust LDS, one microseta between LDS. [Note: The right LDT bore 3 full-sized LDS.]
A2A3: LT with seven to eight long, robust LS, four shorter, straight setae basally. Dorsum with anterior row
of seven to ten (A2, A3) SMS, mesal row of four (A2, A3), posterior row of eight (A2, A3) between LDTs. A4A5:
LT with nine to ten (A4), six (A5) robust, hooked or straight LS apically, dorsally, one to four short, straight LS
basally. LDS long, robust, hooked. Dorsum with anterior row of eight (A4, A5) long SMS, mesal row of four SMS,
posterior row of eleven to eight SMS (A4, A5) between LDTs. A6: Anterior row of three intermediate-length SMS,
no other SMS. A7: LDT with one long, robust, straight LDS, two short, straight LDS. Two to three pairs of very
small setae between spiracles. A8: Row of four short, robust setae between spiracles. Posterior margin with four
short, robust setae.
Biological notes. Like L. (L.) nigrilabris females, L. (L.) insularis females lay their eggs in clusters at the ends
of long, intertwined stalks [Mark S. Fox, personal communication; his photos here (Fig. 21) and on http://
bugguide.net/node/view/350432)].

118 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


L. (L.) insularis larvae have been found in leaf litter, and their debris packets consist largely of small snail
shells, and sometimes live snails. Jones (1929, 1941) reported the following snail species from specimens collected
in Vinton and Washington Counties, Ohio: Punctum vitreum (H. B. Baker) [as Punctum pygmaeum (Drap.)],
Euconulus fulvus (Mller), Striatura milium (Morse), Carychium exiguum (Say), Strobilops labyrinthica (Say),
Cochlicopa lubrica (Mller). Archer (1938) reported the following snail species from larvae that are almost
certainly L. (L.) insularis, in North Carolina and Alabama: Glyphyalinia indentata (Say) [as Retinella indentata
paucilirata], Glyphyalinia carolinensis (Cockerell) [as R. carolinensis wetherbyi], juveniles of Inflectarius rugeli
(Shuttleworth) [as Polygyra rugeli], Hawaiia minuscula (A. Binney), Guppya sterkii (Dall) [as Euconulus sterkii],
Euconulus chersinus (Say), and Vertigo gouldii (A. Binney). The third instar (described here, collected in Swain
County, North Carolina) also carried mainly small snail shells in its packet (Fig. 22); D. C. Dourson (personal
communication, 2005) identified them as Carychium clappi Hubricht, Gastrocopta contracta (Say), G. pentadon
(Say), Punctum blandianum Pilsbry, P. minutissimum (I. Lea), and P. vitreum. For systematic and biological
information on many of the snails listed above, see Dourson & Dourson (2006).

FIGURE 14. Leucochrysa (L.) insularis, variation in head and thoracic coloration. A-C. Head, thorax, dorsal; D-G. Head,
frontal [note the polymorphic coloration of the mesoscutellum (Figs A-C, partially obscured by pins).] [A, D. Jefferson County,
FL, female (prob); B, C, E, G. Dominican Republic (B, E, G, females; C, male); F. State of Veracruz, Mexico, female].

In addition to snail shells, body parts from a diverse array of insects also are reported from the packets. Jones
(1941) described the behavior involved in applying the snail shells to the larval dorsum. He observed that the shells
were held to the larval setae via silken strands; he suggested that the strands originate from a fluid that the tip of the
abdomen applies to the shell and that hardens on the snail shell. He also stated that larvae feed on snails and that
sometimes larvae carry living snails in the packet which are later removed and eaten. Finally, Jones (1941)
mentioned that L. (L.) insularis larvae overwinter hidden within coiled leaves.
Specimens examined (adults and larvae, in addition to the type). UNITED STATES. GA: Richmond Co.,
Augusta, 24.v.1968 (1, AMNH); KY: Carter Co., along Little Sandy River below Grayson Dam (US-CI),
10.viii.1995, B. C. Kondratieff & R. F. Kirchner (1, COSU); LA: St. Tammany Parish, Honey Island Swamp, M.
Fox, 26.x.2009 () (1, cluster of hatched eggs laid in lab, 6 neonates, unfed; UCB); MI: Hinds Co., Jackson,

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 119


24.viii.1968, B. Mather (1, prob., AMNH); NC: Swain Co., Great Smoky Mountains National Park, Purchase
Knob Survey / Site #28, Sap Tree 00ZING 9-6-05, D. Dourson (1L3, UCB, on loan from Great Smoky Mountains
National Park); VA: Federick Co. Rt. 50, Back Cr., 16.vii.1980, Kondratieff (1, COSU). MEXICO. Veracruz:
Crdoba, 11.x.1966, A. & B. Lan (1, ROM). DOMINICAN REPUBLIC. Azua: East side of crest, Sierra Martin
Garcia, 7 km WNW Barrero, k18-21N, 7050W, 860m, 2526.vii.1992, cloud forest adjacent to disturbed forest,
C. Young, R. Davidson, S. Thompson, J. Rawlins (6, 8, CMNH); La Vega: Jarabaco, 4000, ex vegetation, J.
Doyen (1, prob., UCB); San Juan: Estrelleta, Rio Limpio, 650 m, 15.viii.1980, A. Norrbom (1, CMNH);
Santiago: Cordillera Septentrional, Pico Diego de Ocampo, 1.1 km E. summit, 19.35.06N, 70.44.23M, 958 m,
31.v.2004, J. Rawlins, C. Young, C. Nunez, J. Fetzner (1, CMNH); Santiago Rodriguez: 7 km from Moncion, nr.
Mao River, 23.x.1986, R. Greenfield (1?, BPBM); Province unknown: 20.ii.1978, S. A. Marshall (1, DEBU).
UNKNOWN. Sanchez, xii.1915, ac. 4806 (1, 1, AMNH).

FIGURE 15. Leucochrysa (L.) insularis, wings, body, dorsal [Santiago, Cuba, male].

Discussion

1. Are the larval features of L. (L.) nigrilabris and L. (L.) insularis consistent with those of other species in the
subgenus Leucochrysa (Leucochrysa)?
Larvae of L. (L.) nigrilabris and L. (L.) insularis express all of the twelve features previously proposed to
characterize the genus Leucochrysa, except two (# 6 and # 7 below, marked with asterisks). This list, which
initially was derived from two Leucochrysa (Leucochrysa) species and nine Leucochrysa (Nodita) species (see
Adams 1987, Tauber 2004, Mantoanelli et al. 2006, 2011, Tauber et al. 2011), now is generally supported by a
broader diversity of Leucochrysa (Leucochrysa) species. We suspect that these two exceptions, especially # 6 are
the result of reversals; thus, we temporarily retain the two features (# 6 and # 7) on the list of Leucochrysa
characteristics until larvae from more species have been studied.

120 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


FIGURE 16. Leucochrysa (L.) insularis, male abdomen and genitalia. A. Abdomen, lateral; B. Terminal abdominal segments,
lateral; C. Callus cerci; D. Hypandrium internum; E. Gonarcus, lateral; F. Gonarcus, dorsal; G. Gonarcus, dorsofrontal, with
sclerotized plates and connecting semi-sclerotized membrane bearing gonocristae; H. Gonarcus, dorsal, with sclerotized plates
and connecting semi-sclerotized membrane bearing gonocristae [note two patches of gonosetae on membrane below
mediuncus] [A, B, G, Cuba; C-F, H, Dominican Republic]. Abbreviations: bk, beak-like tip of mediuncus; c, comes; c.c., callus
cerci; d.a., dorsal apodeme of T9+ect; fl, mediuncal flange; gs, gonarcus; gse, gonosetae on small chalazae; g.a., gonarcal
apodeme; g.b., gonarcal bridge; m, membranous connection between sclerotized plates, with gonocristae; mu, mediuncus
(fused rods and beak); s.p., sclerotized plate on membrane above tip of S8+9; S8+9, fused eighth and ninth sternites; tr,
trichobothria; T7, seventh tergite; T9+ect, fused ninth tergite and ectoproct.

Larval characteristics of the genus Leucochrysa


1 Thoracic lateral tubercles long: prothoracic tubercles extending at least to mid-head, and meso-and
metathoracic tubercles longer than half the width of the corresponding segment (all instars),
2 Thoracic tubercles with very long setae (all instars), extending in a fan-shape from the apical and lateral
surfaces (L2, L3),
3 Anterior segments of the abdomen thicker than those of the thorax, giving the abdomen a humped appearance
in lateral view (all instars),
4 Abdominal segments A7 to A10 small, curved ventrally, and partially telescoped into the anterior segment (all
instars),

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5 Terminal seta of the antenna at least half the length of the flagellum (all instars),
6 *Prothorax with primary setae S1, S3, S4, and S5 present, S2 absent [all instars, except L2 and L3 of L. (L.)
nigrilabris and L. (L.) insularis],
7 *Mesonotum and metanotum each with two transverse rows of long, smooth, usually hooked setae arising
from chalazae [all instars, except L. (L.) varia, L. (L.) nigrilabris and L. (L.) insularis (see Mantoanelli et al.
2006, descriptions above)],
8 Lateral tubercles on abdominal segments A2 and A3 papilliform, with long setae extending from the anterior,
dorsal and apical surfaces (L2, L3),
9 Lateral tubercles on abdominal segments A4 to A7 slightly elongated, bearing long setae mainly on the apical
surface (L2, L3),
10 Abdominal segments A1 to A5 each with a pair of laterodorsal tubercles, each tubercle with two long, smooth,
hooked setae, separated by microseta [all instars, except microseta sometimes absent from L1: e.g,
Leucochrysa (Nodita) azevedoi Navs (Mantoanelli et al. 2011)],
11 Abdominal segment A1 with a single row of smooth, hooked submedian setae, between the laterodorsal
tubercles (all instars; L1: four setae in row; L2, L3: number of setae variable),
12 Abdominal segments A2 to A5 with two (L1) or three (L2, L3) rows of smooth, hooked submedian setae, the
posterior row between the laterodorsal tubercles (L1: four setae in anterior row, two in posterior row, all
between LDTs; L2, L3: number of setae variable).

FIGURE 17. Leucochrysa (L.) insularis female abdomen. A. Fourth to terminal segments, lateral; B. Sixth to terminal
segments, lateral; C. Callus cerci; D. Subgenitale, lateral, with genitalia removed [A, B. Dominican Republic, mature
specimens; C. Dominican Republic, slightly teneral specimen; D. Veracruz, Mexico, teneral specimen]. Abbreviations: g.l.,
gonapophysis lateralis; sg, subgenitale; sg-bl, bilobed, dorsal process of subgenitale; sg-v, protruding ventral process of
subgenitale; sp, spermatheca; sp.d., spermathecal duct; S7, seventh sternite; tr, trichobothria; T5, T6, fifth and sixth tergites;
T9+ect, fused ninth tergite and ectoproct.

122 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


FIGURE 18. Leucochrysa (L.) insularis female genitalia. A. Spermatheca, lateral; B. Spermatheca, dorsal; C. Dissected
genitalia, bursa copulatrix (torn); D. Colleterial complex, lateral [all, Veracruz, Mexico]. Abbreviations: acc.gl., colleterial
accessory gland; b.c., bursa copulatrix; b.d., bursal duct; b.gl., bursal gland; col.gl., colleterial gland; col. r., colleterial
reservoir; g.l., gonapophysis lateralis; inv, spermathecal invagination; slit, dorsal slit in spermathecal vellum that opens to bursa
copulatrix; sp, spermatheca; sp.d., spermathecal duct; tr.s., transverse sclerite; vel, velum.

Larval characteristics of the subgenus Leucochrysa


Only two larval features are known to differentiate the larvae of subgenus Leucochrysa from those in subgenus
Nodita; both are derived from the spiracles (Mantoanelli et al. 2006, Tauber et al. 2011). For subgenus
Leucochrysa the character states are: (i) spiracles borne on top of plump protuberances (swellings), especially on
the mesothorax and A1, and (ii) spiracular openings surrounded by well-sclerotized rings that project substantially
above the surface of the integument. As with the two species in the subgenus that were previously described in
detail, L. (L.) nigrilabris and L. (L.) insularis larvae express the above attributes.

(2) Do adult and larval features support the synonymy and/or proposed close relationship between the two
species?
A. The findings of this study do not support Taubers (2004) suggestion that L. (L.) nigrilabris might be
synonymous with L. (L.) insularis. Rather, they provide morphological evidence from both adults and larvae that
clearly indicates the two species are distinct.

Differences between L. (L.) nigrilabris and L. (L.) insularis


Male: (i) The gonarcus of L. (L.) nigrilabris, but not L. (L.) insularis, has a pair of gonocornua (Compare Figs
19A and B; also Figs 4AC, E and 16EH). (ii) The gonosetae of L. (L.) nigrilabris are located on a stiff membrane
above the mediuncus, whereas in L. (L.) insularis the gonosetae are on a loosely attached membrane below the

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mediuncus (Figs 3BE, 16H). (iii) The sclerotized genital plates are attached to the S9 in L. (L.) nigrilabris,
whereas in L. (L.) insularis they are attached to each other, but not to S9, by a lightly sclerotized membranous
connection bearing gonocristae (Figs 3B, F, 16G).

FIGURE 19. Species-specific genital characteristics of Leucochrysa (L.) insularis (left column) and L. (L.) nigrilabris (right
column). A, B. Gonarcal complex; C, D. Subgenitale [A, C. Dominican Republic; B, D. Aragua, Venezuela]. Abbreviations:
bk, beak-like tip of mediuncus; gc, gonocornu; g.a., gonarcal apodeme; g.b., gonarcal bridge; mu, mediuncus (fused rods and
beak); sg-bl, bilobed, dorsal process of subgenitale; sg-v, protruding ventral process of subgenitale; sp, spermatheca.

Female: (i) Both species have subgenitale with two posterior processes (lobe of Tauber 2004), the ventral
one is knob-like and withdrawn in L. (L.) nigrilabris, whereas in L. (L.) insularis, it is cylindrical and protruding
posteriorly (Compare Figs 19C and D; also Figs 7B and 17A, B, D) (the dorsal process is bilobed and protruding in
both species). (ii) The L. (L.) nigrilabris spermatheca has rounded lateral walls and a deep, narrow invagination
(Figs 7B, C, 19D); in contrast, the L. (L.) insularis spermatheca has straight, steep lateral walls and a deep, broad
invagination (Figs 18A, B, 19C). (iii). The L. (L.) insularis callus cerci is round as in L. (L.) nigrilabris, but its
trichobothria are larger and fewer (Figs 7, 16, 17).
Larvae: Leucochrysa (L.) nigrilabris and L. (L.) insularis differ in their (i) genal markings (Figs 12G, 20C),
(ii) the robustness of their primary setae, and (iii) the number of thoracic and abdominal setae (e.g., Table 1).

124 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


FIGURE 20. Leucochrysa (L.) insularis third instar. A. Head, dorsal; B. Head, ventral; C. Head, lateral; D. Head and
prothorax; E. Mesothorax, metathorax, and abdominal segments A1A5; F. Abdominal segments 110, dorsal [all, North
Carolina, USA]. Abbreviations: Ax, abdominal segment number; T1LT, prothoracic lateral tubercle; T3, metathorax.

FIGURE 21. Leucochrysa (L.) insularis cluster of hatched eggs; stalk length ~1 cm [Honey Island Swamp, St. Tammany
Parish, LA]. Photo by Mark S. Fox.

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FIGURE 22. Leucochrysa (L.) insularis third instar with debris packet containing snails. A. Dorsal; B. Lateral. [Great Smoky
Mountains National Park, Swain Co., NC]. Photo by Daniel C. Dourson.

TABLE 1. Comparisons of selected larval features among four Leucochrysa (Leucochrysa) species.

L. (L.) varia L. (L.) nigrilabris L. (L.) insularis L. (L.) boxi

Third Instars
Ratio
Mandible length : head width 1.11.2 1.2 1.1 1.4
T1LT length : width 7.07.6 9.510.9 8.3 13.213.5
T1LT length : mandible length 0.50.6 0.7 0.6 0.9
T1LT length : prothoracic length 0.8 1.21.3 -- 1.2

Number of setae
Mesothorax, posterior row 6 1114 4 5660
Metathorax, posterior row 8 1516 12 48
Dorsum, A1 row of SMS 9 10 10 5+18 (2 rows)
A2, posterior row of SMS 67 10 8 12
A3, posterior row of SMS 8 10 8 12

Second Instars
Number of setae
Mesothorax, posterior row 6 5 -- 1618
Metathorax, posterior row 8 11 -- 1618
Dorsum, A1 row of SMS 5 8 -- 810 (1 row)
A2, posterior row of SMS 24 4 -- 68
A3, posterior row of SMS 24 4 -- 68

Note: The values are approximations, based on two to three sets of measurements per value for each species except L. (L.)
insularis for which only one third instar (teneral) was available. The SMS lie between the laterodorsal tubercles and do not
include the LDS.

B. Adult and larval morphology support the hypothesis that L. (L.) nigrilabris and L. (L.) insularis are very closely
related.

Similarities between L. (L.) nigrilabris and L. (L.) insularis


Adult coloration: (i) Southern Caribbean populations of L. (L.) insularis are similar to those of L. (L.)
nigrilabris in the depth of their coloration and body markings, as well as in the pattern of their markings, including,

126 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


in some specimens, the darkened surface of the labrum (compare Figs 1 and 14). (ii) Populations of both species
exhibit a similar type of polymorphism (intrapopulation variation) in the coloration of the mesoscutellum; in both
species, the sclerite can be entirely dark brown, tan or white [L. (L.) nigrilabris: Figs 1DF; L. (L.) insularis: Figs
14AC]. (iii) L. (L.) nigrilabris and L. (L.) insularis both exhibit large dark spots on the fifth and sixth abdominal
tergites.
Abdomen: Both species have round callus cerci (Figs 3B, E, 5A, 6AD, 16A, B, 17AC)]; in many
leucochrysine species they are oval-shaped.
Male: Some characteristics of the male genitalia that are shared by the two species appear to be unique among
members of the subgenus. They are: (i) a narrow, elongate mediuncus that terminates in a relatively large, narrow,
deeply curved hook (Figs 4B, E, 16E, F, H), (ii) gonosaccus with relatively robust gonosetae arising from chalazae
(Figs 4BE, 16F, H), and (iii) terminus of S9 with a pair of flat, sclerotized lobes. The lobes are well developed and
attached to S9 in L. (L.) nigrilabris (Figs 3B, F); they are less developed and located on a partially sclerotized
membranous fold in L. (L.) insularis (Figs 16G, H).
Female: Both species have (i) a simple, doughnut-shaped, invaginated spermatheca, and (ii) either a very
small, simple bursal duct or none at all (Figs 7BD, 18A, B, 19C, D). The spermatheca opens to the bursa
copulatrix via a slit in the spermathecal velum (Figs 7D, 18B), and the bursal glands are elongate, tubular and
unbranched (Fig. 7D, 18C). The above two features are not unique among females of the subgenus [e.g., the
spermathecal characteristics occur in Leucochrysa (L.) colombia Banks (Adams 1977) and L. (L.) boxi (Tauber et
al. 2011)]; nevertheless, they separate L. (L.) nigrilabris and L. (L.) insularis from the Leucochrysa (Leucochrysa)
varia-like species, all of which have elongate, tubular, often coiled spermathecae, well developed bursal ducts,
and bulbous bursal glands.
On the basis of the above shared, distinguishing male and female features, we consider that L. (L.) nigrilabris
and L. (L.) insularis are very closely related to each other, but not to the varia-like species.
Larvae: Leucochrysa (L.) nigrilabris and L. (L.) insularis share some features not yet reported for other
Leucochrysa (Leucochrysa) species (Table 1). Both have: (i) similar ratios of prothoracic lateral tubercle length :
width (= 8.710.9, slightly longer than that of L. (L.) varia, but shorter than that of L. (L.) boxi)], and (ii) S2 present
on the prothorax of L2 and L3.
Biology: Both L. (L.) nigrilabris and L. (L.) insularis females lay their eggs in clusters at the ends of long,
intertwined stalks (Fig. 21). This type of oviposition behavior has been reported for a few species in the chrysopine
genera Pseudomallada, Nineta, and Ungla (Withycombe 1922, Principi 1940, 1956, Duelli 1984, Gepp 1988,
Daz-Aranda & Monserrat 1991, Tsukaguchi 1995, Freitas 2003), but L. (L.) nigrilabris and L. (L.) insularis have
the only records for the Leucochrysini.
It is noteworthy that L. (L.) insularis larvae are found in the leaf litter, and that they prey on small snails, whose
shells they place and carry on their dorsa. We have not observed L. (L.) nigrilabris larvae in the field and thus do
not know whether they resemble those of L. (L.) insularis in their habitat association, feeding habits, or trash
packets.

(3) Should the two species be aligned with the L. (L.) varia-like species?
Although L. (L.) nigrilabris and L. (L.) insularis adults share a number of external features with the L. (L.)
varia-like species, the results from our studies here and elsewhere (Tauber et al. 2013) do not provide support for
a particularly close relationship among these taxa.
Adults: The most obvious shared adult characteristics of the three species include (i) a conspicuous
polymorphism in mesothoracic markings, (ii) red to reddish brown scapes, (iii) vertex with red to reddish brown
lateral marks, and (iv) dark shading around the distal Psm and Psc crossveins of the forewing (Figs 1, 2, 14, 15).
In contrast, the L. (L.) nigrilabris and L. (L.) insularis adults lack many other external features of the varia-
like species (see Tauber et al. 2013); i.e., they do not have the (i) red coloration on the frons, gena and scapes or
(ii) shading around the second m-cu2 crossvein of the forewing that the varia-like species usually express. More
importantly, their genital characteristics are distinct from those of the varia-like species. For example, in L. (L.)
nigrilabris and L. (L.) insularis females, (iii) the spermatheca is simple, and it lacks a coiled, tubular bursal duct
that typifies the varia-like species. In the males, (iv) the mediuncus is narrow and elongate, and it terminates in a
slender, elongate and deeply curved mediuncal beak (c.f. Tauber et al. 2013). Thus, on the basis of adult features, it
appears that the L. (L.) nigrilabris and L. (L.) insularis should not be grouped with the varia-like species.

LEUCOCHRYSA ( LEUCOCHRYSA ) Zootaxa 3750 (2) 2013 Magnolia Press 127


Larvae: The situation is less clear for the larvae. Of the four Leucochrysa (Leucochrysa) species that have
detailed larval descriptions, L. (L.) nigrilabris, L. (L.) insularis, and L. (L.) varia share a number of features that
distinguish them from L. (L.) boxi (Mantoanelli et al. 2006, Tauber et al. 2011). For example, (i) their head
markings, which are very similar, differ from those of L. (L.) boxi; (ii) they have similar ratios of mandible length :
head width that are shorter than those of L. (L.) boxi, (iii) their mesothorax and metathorax have only one
transverse row of hooked setae [not two rows as in L. (L.) boxi], and (iv) they have fewer setae overall than does L.
(L.) boxi (Figs 12A, 21A, Table 1).
However, L. (L.) nigrilabris and L. (L.) insularis (L2, L3) differ markedly from L. (L.) varia in a number of
other important traits: (i) their thoracic lateral tubercles are longer (relative to their width and relative to the length
of the prothorax) (See Fig. 9, Table 1); (ii) the head markings, mandibles and setae are darker brown; (iii) the dorsal
setae are larger, more robust, and, on some segments, more numerous (Table 1); and (iv) the posterior sections of
the mesothorax and metathorax have more setae than those of L. (L.) varia.
The first instars of L. (L.) nigrilabris, L. (L.) insularis, and L. (L.) varia have very similar setal patterns;
however, there are some important exceptions. For example, in L. (L.) nigrilabris and L. (L.) insularis, the
prothoracic lateral tubercle (LT) and the abdominal LDTs lack a microseta between the two long setae (LS, LDS)
on the tubercles, whereas in the L. (L.) varia first instars, those microsetae are present on both types of tubercles.

Conclusion

On the basis of both adult and larval characters, it appears that L. (L.) nigrilabris and L. (L.) insularis are very
closely related to each other. In turn, these two species express several adult (external) and larval features that are
consistent with those of L. (L.) varia; these similarities may indicate a relationship between the two sets of species.
However, the genitalia (male & female) of L. (L.) nigrilabris and L. (L.) insularis differ significantly from those of
the L. (L.) varia-like species. Thus, at this time, we refrain from grouping the two sets of species together.

Acknowledgements

We thank the following individuals for facilitating our study: J. K. Liebherr & E. R. Hoebeke (CUIC), P. Perkins, S.
P. Cover & B. D. Farrell (MCZ), C. B. Barr & P. Oboyski (UCB), D. Goodger (BMNH), J. E. Rawlins (CMNH), O.
S. Flint, D. G. Furth & T. Erwin (USNM), S. A. Marshall (DEBU), S. Heyden & L. S. Kimsey (UCD), D. C.
Darling (ROM), W. Xie & B. V. Brown (LACM), B. Kondratieff (COSU), J. Demarmels, J. Clavijo & Q. rias
(MIZA), N. W. Perioto & R.I.R. Lara, Agncia Paulista de Tecnologia dos Agronegcios, D. A. Grimaldi & T.
Nguyen (AMNH), N. L. Evenhuis & S. Meyers (BPBM).
We also thank E. A. Silva, A. J. Tauber, and P. J. Tauber, who assisted with collecting specimens; E.
Mantoanelli, who helped with larval rearing; C. R. Parker (USGS, NPS, Great Smoky Mountains NP), P. E. Super
(NPS, Great Smoky Mountains NP), & R. Nichols (NPS, Great Smoky Mountains NP) for facilitating our study of
the L. (L.) insularis larva; D. C. Dourson (Lincoln Memorial University, Harrogate, TN) for use of his photos of the
L. (L.) insularis larva, for providing the specimen, and for his help with the snail identifications; and M. S. Fox
(Tulane University, New Orleans) for use of his photo of the L. (L.) insularis egg cluster and for sending the
voucher specimens. The Lacewing Digital Library was helpful during this study and we acknowledge J. D.
Oswald (Texas A & M University, College Station, TX) for access.
This work received support from the National Science Foundation (Grants INT-9817231, DEB-0542373, MJT,
CAT), the National Geographic Society (MJT, CAT, GSA), the USDA/NRI (Competitive Grant 9802447, MJT,
CAT), Conselho Nacional de Desenvolvimento Cientfico e Tecnolgico (CNPq, Brazil Grants 300504/96-9,
466439/00-8, 475848/04-7, 484497/07-3, GSA), Regional Project W-3185, Cornell University, and the
Universidade Estadual do Norte Fluminense.

128 Zootaxa 3750 (2) 2013 Magnolia Press TAUBER ET AL.


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